Tonotopic Place Research Articles

Cochlear microphonic latency predicts outer hair cell function in animal models and clinical populations

As recently reported, electrocochleography recorded in cochlear implant recipients showed reduced amplitude and shorter latency in patients with more severe high-frequency hearing loss compared with those with some residual hearing. As the response is generated primarily by receptor currents in outer hair cells, these variations in amplitude and latency may indicate outer hair cell function after cochlear implantation. We propose that an absence of latency shift when the cochlear microphonic is measured on two adjacent electrodes indicates an absence or dysfunction of outer hair cells between these electrodes. We test this preclinically in noise deafened guinea pigs (2 h of a 124 dB HL, 16–24 kHz narrow-band noise), and clinically, in electrocochleographic recordings made in cochlear implant recipients immediately after implantation. We found that normal hearing guinea pigs showed a progressive increase in latency from basal to apical electrodes. In contrast, guinea pigs with significantly elevated high-frequency hearing thresholds showed no change in cochlear microphonic latency measured on basal electrodes (located approximately at the 16–24 kHz location in the cochlea).. In the clinical cohort, a significant negative correlation existed between cochlear microphonic latency shifts and hearing thresholds at 1-, 2-, & 4 kHz when tested on electrodes located at the relevant cochlear tonotopic place. This reduction in latency shift was such that patients with no measurable hearing also had no detectable latency shift (place assessed by CT scan, r’s of -.70 to -.83). These findings suggest that electrocochleography can be used as a diagnostic tool to detect cochlear regions with functioning hair cells, which may be important for defining cross-over point for electro-acoustic stimulation.

Nonlinear reflection as a cause of the short-latency component in stimulus-frequency otoacoustic emissions simulated by the methods of compression and suppression.

Stimulus-frequency otoacoustic emissions (SFOAEs) are generated by coherent reflection of forward traveling waves by perturbations along the basilar membrane. The strongest wavelets are backscattered near the place where the traveling wave reaches its maximal amplitude (tonotopic place). Therefore, the SFOAE group delay might be expected to be twice the group delay estimated in the cochlear filters. However, experimental data have yielded steady-state SFOAE components with near-zero latency. A cochlear model is used to show that short-latency SFOAE components can be generated due to nonlinear reflection of the compressor or suppressor tones used in SFOAE measurements. The simulations indicate that suppressors produce more pronounced short-latency components than compressors. The existence of nonlinear reflection components due to suppressors can also explain why SFOAEs can still be detected when suppressors are presented more than half an octave above the probe-tone frequency. Simulations of the SFOAE suppression tuning curves showed that phase changes in the SFOAE residual as the suppressor frequency increases are mostly determined by phase changes of the nonlinear reflection component.

Using Cochlear Microphonic Potentials to Localize Peripheral Hearing Loss.

The cochlear microphonic (CM) is created primarily by the receptor currents of outer hair cells (OHCs) and may therefore be useful for identifying cochlear regions with impaired OHCs. However, the CM measured across the frequency range with round-window or ear-canal electrodes lacks place-specificity as it is dominated by cellular sources located most proximal to the recording site (e.g., at the cochlear base). To overcome this limitation, we extract the “residual” CM (rCM), defined as the complex difference between the CM measured with and without an additional tone (saturating tone, ST). If the ST saturates receptor currents near the peak of its excitation pattern, then the rCM should reflect the activity of OHCs in that region. To test this idea, we measured round-window CMs in chinchillas in response to low-level probe tones presented alone or with an ST ranging from 1 to 2.6 times the probe frequency. CMs were measured both before and after inducing a local impairment in cochlear function (a 4-kHz notch-type acoustic trauma). Following the acoustic trauma, little change was observed in the probe-alone CM. In contrast, rCMs were reduced in a frequency-specific manner. When shifts in rCM levels were plotted vs. the ST frequency, they matched well the frequency range of shifts in neural thresholds. These results suggest that rCMs originate near the cochlear place tuned to the ST frequency and thus can be used to assess OHC function in that region. Our interpretation of the data is supported by predictions of a simple phenomenological model of CM generation and two-tone interactions. The model indicates that the sensitivity of rCM to acoustic trauma is governed by changes in cochlear response at the ST tonotopic place rather than at the probe place. The model also suggests that a combination of CM and rCM measurements could be used to assess both the site and etiology of sensory hearing loss in clinical applications.

Basal contributions to short-latency transient-evoked otoacoustic emission components.

The presence of short-latency (SL), less compressive-growing components in bandpass-filtered transient-evoked otoacoustic emission (TEOAE) waveforms may implicate contributions from cochlear regions basal to the tonotopic place. Recent empirical work suggests a region of SL generation between ∼1/5 and 1/10-octave basal to the TEOAE frequency's tonotopic place. However, this estimate may be biased to regions closer to the tonotopic place as the TEOAE extraction technique precluded measurement of components with latencies shorter than ∼5 ms. Using a variant of the non-linear, double-evoked extraction paradigm that permitted extraction of components with latencies as early as 1 ms, the current study empirically estimated the spatial-extent of the cochlear region contributing to 2 kHz SL TEOAE components. TEOAEs were evoked during simultaneous presentation of a suppressor stimulus, in order to suppress contributions to the TEOAE from different places along the cochlear partition. Three or four different-latency components of similar frequency content (∼2 kHz) were identified for most subjects. Component latencies ranged from 1.4 to 9.6 ms; latency was predictive of the component's growth rate and the suppressor frequency to which the component's magnitude was most sensitive to change. As component latency decreased, growth became less compressive and suppressor-frequency sensitivity shifted to higher frequencies. The shortest-latency components were most sensitive to suppressors approximately 3/5-octave higher than their nominal frequency of 2 kHz. These results are consistent with a distributed region of generation extending to approximately 3/5-octave basal to the TEOAE frequency's tonotopic place. The empirical estimates of TEOAE generation are similar to model-based estimates where generation of the different-latency components occurs through linear reflection from impedance discontinuities distributed across the cochlear partition.

The effect of stimulus bandwidth on the nonlinear-derived tone-burst-evoked otoacoustic emission.

Intermodulation distortion has been hypothesized as a mechanism contributing to the generation of short-latency (SL) components in the transient-evoked otoacoustic emission (TEOAE). Presumably, nonlinear interactions between the frequency components within the evoking stimulus induce cochlear distortion products, which mix in the cochlea and ear canal with reflected energy from each stimulus-frequency's tonotopic place. The mixing of these different components is evidenced in the bandpass-filtered emission waveform as a series of different latency peaks. The current study tested the hypothesis that intermodulation distortion, induced within the spectral bandwidth of the evoking stimulus, is the primary mechanism through which the SL components are generated. The nonlinear-derived tone-burst-evoked OAE (TBOAEnl) was evoked using 2-kHz tone bursts with durations of 3, 6, 12, and 24 cycles. As tone burst duration doubled, the spectral bandwidth was halved. It was hypothesized that contributions to the TBOAEnl from SL components would decrease as tone burst duration increased and spectral bandwidth decreased, if the SL components were generated through intermodulation distortion. Despite differences in spectral bandwidth between the evoking stimuli, the latencies and magnitudes of the different latency components between the 3- and 6-cycle TBOAEnl were comparable. The 12- and 24-cycle TBOAEnl envelopes were characteristic of destructive phase interactions between different latency components overlapping in time. The different latency components in the 3- and 6-cycle TBOAEnl introduced a characteristic level dependency to TBOAEnl magnitude and latency when analyzed across a broad time window spanning the different components. A similar dependency described the 12- and 24-cycle TBOAEnl input/output and latency-intensity functions, suggesting that the SL components evident in the shorter-duration TBOAEnl equally contributed to the longer-duration TBOAEnl, despite reductions in spectral bandwidth. The similarity between the different TBOAEnl suggests that they share a common generation mechanism and casts doubt on intermodulation distortion as the generation mechanism of SL TEOAE components in humans.

Estimating cochlear frequency selectivity with stimulus-frequency otoacoustic emissions in chinchillas.

It has been suggested that the tuning of the cochlear filters can be derived from measures of otoacoustic emissions (OAEs). Two approaches have been proposed to estimate cochlear frequency selectivity using OAEs evoked with a single tone (stimulus-frequency (SF)) OAEs: based on SFOAE group delays (SF-GDs) and on SFOAE suppression tuning curves (SF-STCs). The aim of this study was to evaluate whether either SF-GDs or SF-STCs obtained with low probe levels (30 dB SPL) correlate with more direct measures of cochlear tuning (compound action potential suppression tuning curves (CAP-STCs)) in chinchillas. The SFOAE-based estimates of tuning covaried with CAP-STCs tuning for >3 kHz probe frequencies, indicating that these measures are related to cochlear frequency selectivity. However, the relationship may be too weak to predict tuning with either SFOAE method in an individual. The SF-GD prediction of tuning was sharper than CAP-STC tuning. On the other hand, SF-STCs were consistently broader than CAP-STCs implying that SFOAEs may have less restricted region of generation in the cochlea than CAPs. Inclusion of <3 kHz data in a statistical model resulted in no significant or borderline significant covariation among the three methods: neither SFOAE test appears to reliably estimate an individual's CAP-STC tuning at low-frequencies. At the group level, SF-GDs and CAP-STCs showed similar tuning at low frequencies, while SF-STCs were over five times broader than the CAP-STCs indicating that low-frequency SFOAE may originate over a very broad region of the cochlea extending ≥5 mm basal to the tonotopic place of the probe.

Plasticity in human pitch perception induced by tonotopically mismatched electro-acoustic stimulation

Under normal conditions, the acoustic pitch percept of a pure tone is determined mainly by the tonotopic place of the stimulation along the cochlea. Unlike acoustic stimulation, electric stimulation of a cochlear implant (CI) allows for the direct manipulation of the place of stimulation in human subjects. CI sound processors analyze the range of frequencies needed for speech perception and allocate portions of this range to the small number of electrodes distributed in the cochlea. Because the allocation is assigned independently of the original resonant frequency of the basilar membrane associated with the location of each electrode, CI users who have access to residual hearing in either or both ears often have tonotopic mismatches between the acoustic and electric stimulation. Here we demonstrate plasticity of place pitch representations of up to three octaves in Hybrid CI users after experience with combined electro-acoustic stimulation. The pitch percept evoked by single CI electrodes, measured relative to acoustic tones presented to the non-implanted ear, changed over time in directions that reduced the electro-acoustic pitch mismatch introduced by the CI programming. This trend was particularly apparent when the allocations of stimulus frequencies to electrodes were changed over time, with pitch changes even reversing direction in some subjects. These findings show that pitch plasticity can occur more rapidly and on a greater scale in the mature auditory system than previously thought possible. Overall, the results suggest that the adult auditory system can impose perceptual order on disordered arrays of inputs.

Pitch and loudness matching of unmodulated and modulated stimuli in cochlear implantees

The pitch elicited by unmodulated and amplitude modulated electrical pulse trains was examined with six adult cochlear implantees. In addition, for three of those subjects who had some hearing in their contralateral ear, the pitch of unmodulated electrical pulse trains was compared to that of complex harmonic acoustic tones. In the first experiment, pulse rate discrimination and the effects of place and level differences on pitch were examined for unmodulated pulse trains. General results were consistent with previous studies showing that variations in pulse rate, while holding loudness fixed, elicit changes in pitch at low rates, but become progressively harder to discriminate as rates approach approximately 300 pulses-per-second. Variations in place or level of stimulation generally produced changes in pitch consistent with tonotopic place and spread of excitation. In the second experiment, pitch and loudness of unmodulated pulse trains were compared with those of amplitude modulated stimuli as a function of modulation depth, rate, and shape, and presentation level. The pitch elicited by an amplitude modulated pulse train was generally higher than that of an unmodulated pulse train with a pulse rate equal to the modulation rate, and generally decreased toward that of the unmodulated pulse train as modulation depth or rate increased, or as presentation level decreased. Sharper/narrower modulation produced lower pitch. In the final experiment, the pitch heights of acoustic complex harmonic tones and unmodulated pulse trains were compared. When electrical pulse rate was equal to the fundamental frequency of the acoustic tone, similar pitch heights were elicited. The results from these experiments indicate that F0 rate pitch derived from the temporal envelope in existing clinical cochlear implant strategies may often be higher than that of acoustic harmonic tones at the same F0 in normal hearing, and that pitch growth with increasing F0 may be shallower. The relationship between F0 and rate pitch is expected to be more similar to acoustic stimulation for low F0 rates when using new pitch coding strategies that code F0 information via deep (narrow) amplitude modulation of the stimulus envelope. Although that similarity reduces as F0 approaches the upper limit of rate-pitch discrimination, that limit is reached sooner for the shallow (or broad) modulators used in existing clinical strategies.

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Comparison of cochlear delay estimates using otoacoustic emissions and auditory brainstem responses

Different attempts have been made to directly measure frequency specific basilar membrane (BM) delays in animals, e.g., laser velocimetry of BM vibrations and auditory nerve fiber recordings. The present study uses otoacoustic emissions (OAEs) and auditory brainstem responses (ABRs) to estimate BM delay non-invasively in normal-hearing humans. Tone bursts at nine frequencies from 0.5 to 8 kHz served as stimuli, with care taken to quantify possible bias due to the use of tone bursts with different rise times. BM delays are estimated from the ABR latency estimates by subtracting the neural and synaptic delays. This allows a comparison between individual OAE and BM delays over a large frequency range in the same subjects, and offers support to the theory that OAEs are reflected from a tonotopic place and carried back to the cochlear base via a reverse traveling wave.

Two-tone suppression of stimulus frequency otoacoustic emissions in human ears with sensorineural hearing loss

Two-tone suppression (2TS) of stimulus frequency otoacoustic emissions (SFOAEs) recorded in normal-functioning human ears is a noninvasive measure of cochlear nonlinearity. This study investigates 2TS SFOAE response properties in 63 subjects with sensorineural hearing loss and normal middle-ear function. SFOAEs were measured using tones at frequencies (fp) from 0.5 to 8 kHz and levels from 40-60 dB SPL. For interpreting SFOAEs, hearing status was classified at each frequency according to degree of hearing loss, with losses ranging from within normal limits to 95 dB HL. SFOAE suppression was measured as a function of suppressor frequency (fs) and level (Ls), with results analyzed in terms of a SFOAE suppression tuning curve. A tip-to-tail difference (TTD) was defined between the Ls needed to produce a criterion decrement in SFOAE level at fs one octave below fp and the Ls needed to produce a criterion decrement near fp. The TTD estimates cochlear-amplifier gain in normal-functioning ears, but its properties in impaired ears based on SFOAEs have not been examined. Results will be analyzed as a function of the hearing status to assess the relative impact of outer-hair cell dysfunction in the SFOAE source region at tonotopic place. [Work supported by NIH.]

Transient evoked otoacoustic emission latency and cochlear tuning at different stimulus levels

Cochlear latency has been evaluated in young adults by time-frequency analysis of transient evoked otoacoustic emissions recorded using the nonlinear acquisition mode at different levels of the click stimulus. Objective, even if model-dependent, estimates of cochlear tuning have been obtained from the otoacoustic latency estimates. Transmission-line cochlear models predict that the transient-evoked otoacoustic emission latency is dependent on the stimulus level, because the bandwidth of the cochlear filter (tuning) depends on the local cochlear excitation level due to nonlinear damping. The results of this study confirm the increase of tuning with increasing frequency and show clearly the decrease of latency and tuning with increasing stimulus level. This decrease is consistent with the expected relation between the slowing down of the traveling wave near the tonotopic place and the cochlear excitation amplitude predicted by cochlear models including nonlinear damping. More specifically, these results support the models in which nonlinear damping consists of a quadratic term and a constant positive term.

The reflectivity function of the cochlear transmission line

The generation of evoked otoacoustic emissions is related, for each frequency, to the reflectivity function of the cochlear membrane near the correspondent resonant place. A formal analogy exists between the transmission line equation that is often used to model the propagation of sound along the cochlea and the Schroedinger equation describing the motion of an elementary particle in a one-dimensional potential. In this analogy, the cochlear region near the tonotopic place is equivalent to a negative potential well. Analytical solutions are available in quantum mechanics textbooks, which predict partial reflection of the incoming wave from such a potential well. The reflectivity is expected to be a slow quasi-periodic function of the well width and depth, e.g., of the quality factor of the cochlear resonance. This observation could be useful to predict the large-scale spectral structure of evoked otoacoustic emissions as a function of the cochlear tuning curve.

Frequency-place compression and expansion in cochlear implant listeners

In multichannel cochlear implants, low frequency information is delivered to apical cochlear locations while high frequency information is delivered to more basal locations, mimicking the normal acoustic tonotopic organization of the auditory nerves. In clinical practice, little attention has been paid to the distribution of acoustic input across the electrodes of an individual patient that might vary in terms of spacing and absolute tonotopic location. In normal-hearing listeners, Başkent and Shannon (J. Acoust. Soc. Am. 113, 2003) simulated implant signal processing conditions in which the frequency range assigned to the array was systematically made wider or narrower than the simulated stimulation range in the cochlea, resulting in frequency-place compression or expansion, respectively. In general, the best speech recognition was obtained when the input acoustic information was delivered to the matching tonotopic place in the cochlea with least frequency-place distortion. The present study measured phoneme and sentence recognition scores with similar frequency-place manipulations in six Med-El Combi 40+ implant subjects. Stimulation locations were estimated using the Greenwood mapping function based on the estimated electrode insertion depth. Results from frequency-place compression and expansion with implants were similar to simulation results, especially for postlingually deafened subjects, despite the uncertainty in the actual stimulation sites of the auditory nerves. The present study shows that frequency-place mapping is an important factor in implant performance and an individual implant patient's map could be optimized with functional tests using frequency-place manipulations.

Spontaneous Basilar Membrane Oscillation and Otoacoustic Emission at 15 kHz in a Guinea Pig

A spontaneous otoacoustic emission (SOAE) measured in the ear canal of a guinea pig was found to have a counterpart in spontaneous mechanical vibration of the basilar membrane (BM). A spontaneous 15-kHz BM velocity signal was measured from the 18-kHz tonotopic location and had a level close to that evoked by a 14-kHz, 15-dB SPL tone given to the ear. Lower-frequency pure-tone acoustic excitation was found to reduce the spontaneous BM oscillation (SBMO) while higher-frequency sound could entrain the SBMO. Octave-band noise centered near the emission frequency showed an increased narrow-band response in that frequency range. Applied pulses of current enhanced or suppressed the oscillation, depending on polarity of the current. The compound action potential (CAP) audiogram demonstrated a frequency-specific loss at 8 and 12 kHz in this animal. We conclude that a relatively high-frequency spontaneous oscillation of 15 kHz originated near the 15-kHz tonotopic place and appeared at the measured BM location as a mechanical oscillation. The oscillation gave rise to a SOAE in the ear canal. Electric current can modulate level and frequency of the otoacoustic emission in a pattern similar to that for the observed mechanical oscillation of the BM.

Multiple internal reflections in the cochlea and their effect on DPOAE fine structure.

In recent years, evidence has accumulated in support of a two-source model of distortion product otoacoustic emissions (DPOAEs). According to such models DPOAEs recorded in the ear canal are associated with two separate sources of cochlear origin. It is the interference between the contributions from the two sources that gives rise to the DPOAE fine structure (a pseudoperiodic change in DPOAE level or group delay with frequency). Multiple internal reflections between the base of the cochlea (oval window) and the DP tonotopic place can add additional significant components for certain stimulus conditions and thus modify the DPOAE fine structure. DPOAEs, at frequency increments between 4 and 8 Hz, were recorded at fixed f2/f1 ratios of 1.053, 1.065, 1.08, 1.11, 1.14, 1.18, 1.22, 1.26, 1.30, 1.32, 1.34, and 1.36 from four subjects. The resulting patterns of DPOAE amplitude and group delay (the negative of the slope of phase) revealed several previously unreported patterns in addition to the commonly reported log sine variation with frequency. These observed "exotic" patterns are predicted in computational simulations when multiple internal reflections are included. An inverse FFT algorithm was used to convert DPOAE data from the frequency to the "time" domain. Comparison of data in the time and frequency domains confirmed the occurrence of these "exotic" patterns in conjunction with the presence of multiple internal reflections. Multiple internal reflections were observed more commonly for high primary ratios (f2/f1 > or = 1.3). These results indicate that a full interpretation of the DPOAE level and phase (group delay) must include not only the two generation sources, but also multiple internal reflections.

SFOAE input/output functions elicited by slightly off-frequency suppressors

Basilar membrane (BM) response growth is thought to be related to stimulus frequency otoacoustic emission (SFOAE) input/output (I/O) functions. SFOAEs comprise the response to sinusoidal excitation and provide an estimate of nonlinearity at a tonotopic place on the BM. SFOAE I/O functions at frequency f0 (across levels L0) are measured in the presence of suppressor fs (at level Ls) in subjects with normal hearing. fs is varied below, equal to, and above f0 in the range 0.93&amp;lt;fs/f0&amp;lt;1.07. For fixed suppressor level (Ls=80 dB SPL) and varying L0&amp;lt;Ls, the SFOAE I/O function is predicted to be closely related to the BM I/O function. Measured I/O functions are approximately invariant as fs is varied. Functions saturate at increasing levels with a decrease in some ears at the highest levels. An alternative paradigm in which Ls=L0 are covaried yields SFOAE I/O functions with local minima, but which may have a less direct connection to BM response growth. Based on primaries at fs and f0, SFOAE and DPOAE I/O functions differ. One implication is that SFOAE I/O functions may be used to relate BM nonlinear response growth to perceptual processes, including loudness growth and forward masking. [Work supported by NIH (R01 DC003784, T32 DC00013).]

Spectral shapes of forward and reverse transfer functions between ear canal and cochlea estimated using DPOAE input/output functions.

It is widely assumed that the distortion characteristics of the cochlea are uniform across its length, or at least across some portion of its length. For each distortion product otoacoustic emission (DPOAE) input/output (I/O) function across frequency, there is a corresponding cochlear I/O function defined over the cochlear source region. An assumption of distortion invariance is adopted such that these cochlear I/O functions are identical across tonotopic place, which is testable in the sense that a single nonlinear function should adequately describe the set of DPOAE I/O functions across frequency. If so, the differences in measured DPOAE I/O functions across frequency are produced by differences in the forward stimulus transmission to the generation site, and reverse DP transmission back to the ear canal. The absolute transfer-function magnitude is not determined by this technique, but the spectral shapes across frequency and between ears are determined. The role of middle-ear functioning is implicit in the I/O functions because of its controlling influence on these transfer functions. Results have been obtained using the average DPOAE I/O functions measured in a population of healthy ears [Gorga et al., J. Acoust. Soc. Am. 107, 2128-2135 (2000)], and support the hypothesis of cochlear-distortion invariance. The measured forward and reverse transfer functions have a general bandpass characteristic, and a more narrow-band structure with similarities to the behavioral threshold curve.

Model studies of source term and reflection components of synchronous-evoked and distortion product otoacoustic emissions

The unified model of synchronous-evoked, distortion product, and spontaneous otoacoustic emissions of Talmadge et al. [J. Acoust. Soc. Am. 104, 1517 (1998); 105, 275 (1999); 108, 2570 (2000)] provides a useful testing ground for various paradigms that have been proposed for separately determining source terms and reflectance components of synchronous-evoked and distortion-product otoacoustic emissions. The source terms are the ear canal components that would exist in the absence of effects associated with cochlear wave reflections due to, e.g., distributed inhomogeneities [Zweig and Shera, J. Acoust. Soc. Am. 98, 2018 (1995)] and nonlinearity [Talmadge et al., 2000]. Model calculations are used to compare results for the unmixing of these components obtained using reflection-suppression techniques [see, e.g., Kalluri and Shera, J. Acoust. Soc. Am. 109, 622 (2001)] and various forms of time-domain latency windowing ones [see, e.g., Knight and Kemp, J. Acoust. Soc. Am. 107, 457 (2000)]. Because of multiple internal wave reflections at the cochlear base and the tonotopic place for the cochlear waves, the model implies (and experimental data indicate) that the reflectance components will generally exhibit fine structure effects similar to those of the total ear canal signal.

Effects of basilar membrane roughness and nonlinearity on stimulus frequency otoacoustic emissions

Stimulus frequency otoacoustic emissions (SFOAEs) may be evoked via basilar membrane (BM) nonlinearity, BM roughness (random inhomogeneities), or a combination of these properties. Emissions generated by BM nonlinearity alone have phase gradients that are too small to account for the observed emission fine structure [e.g., Shera and Guinan (1999)]. Linear reflection via BM roughness can account for the main phase characteristics of the emissions [e.g., Shera and Zweig (1993); Talmadge et al. (1995)]. It is shown theoretically that, contrary to what is expected from naive considerations concerning phase derivatives and group delays, the physical time delays associated with both emission mechanisms are approximately equal to the round-trip cochlear wave travel time from the cochlear base to the tonotopic place for the stimulus frequency. A theoretical framework is presented for describing the combined effects of roughness and nonlinearity on the emission fine structure. For the case of cochlear roughness and a weak nonlinearity, it is shown that in some instances, the nonlinearity can significantly modify the level and phase behaviors of the fine structure and actually enhance the fine structure patterns. The implications of these results for the stimulus level dependence of SFOAE fine structure are also explored.